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Revista Mexicana de Biodiversidad 81: 863 - 874, 2010
Recibido: 23 febrero 2009; aceptado: 08 enero 2010
Mammalian diversity in climatic domains for Tehuacn-Cuicatln Biosphere
Reserve, Mexico
Diversidad de mamferos en los dominios climticos de la Reserva de la Biosfera
Tehuacn-Cuicatln, Mxico
Oswaldo Tllez Valds1, Vernica Faras1*, Patricia Dvila Aranda1, Janet Louis Stein2, Rafael Lira Saade1
and Francisco J. Botello3
1Laboratorio de Recursos Naturales, Unidad de Biologa, Tecnologa y Prototipos (UBIPRO), Facultad de Estudios Superiores Iztacala Universidad
Nacional Autnoma de Mxico (UNAM). Avenida de los Barrios 1, Los Reyes Iztacala, Tlalnepantla, C.P. 54090, Estado de Mxico, Mxico2Centre for Resource and Environmental Studies, The Australian University, Canberra, Australia.3Laboratorio de Sistema de Informacin Geogrca, Instituto de Biologa, Universidad Nacional Autnoma de Mxico (UNAM). Circuito exterior s/n,
Ciudad Universitaria, Coyoacn, C.P. 04510, Mxico, D. F., Mxico
*Correspondent:[email protected]
Abstract. The Tehuacn-Cuicatln biosphere reserve (BRTC) is rich in mammalian diversity, but geographicaldistribution information is absent or insufcient for most species. Consequently, previous efforts to model the ecological
niche and potential distribution of mammals have been hampered. The main purpose of this study was to examine the
patterns of mammalian diversity in BRTC using a climatic domains classication. Biological datasets composed of
geographically referenced localities commonly are raw input during analyses of geographical distributions of species,
but in countries like Mexico datasets frequently are incomplete and biased. The recent availability of interpolators and
geographic information systems make possible the enhancement of environmental datasets and open the possibility to
use climatic parameters to explain biological patterns. In this study we generated a climatic domain classication for
the Tehuacn-Cuicatln valley and its surrounding areas of inuence. With this approach, climatic domains were used
as biodiversity surrogates, and we justied the overlapping of environmental data with the biological dataset (species,
longitude, latitude, and elevation) to evaluate and complement the available mammal diversity information within
BRTC.
Key words: biodiversity surrogate, geographic distribution, Mammalia.
Resumen. La reserva de la biosfera Tehuacn-Cuicatln (BRTC) posee gran diversidad de mamferos, pero lainformacin sobre distribucin geogrca es incompleta para la mayora de las especies. Esto ha representado una
dicultad en esfuerzos previos para modelar el nicho ecolgico y la distribucin potencial de mamferos en la BRTC.
Nuestro objetivo fue comparar los patrones de diversidad de mamferos en la BRTC usando una clasicacin de dominios
climticos. Las bases de datos biolgicas compuestas de localidades georeferenciadas generalmente son usadas como
datos crudos en anlisis de distribucin geogrca de especies, pero en pases como Mxico frecuentemente estn
incompletas y presentan sesgos de muestreo. La reciente disponibilidad de interpoladores y de sistemas de informacin
geogrca permiti mejorar las bases de datos ambientales, lo que abri la posibilidad de usar parmetros climticos
para explicar patrones biolgicos. En este estudio generamos una clasicacin de dominios climticos del valle de
Tehuacn-Cuicatln y sus reas de inuencia. Con este enfoque, los dominios climticos fueron usados como sustituto
de biodiversidad, y justicamos la superposicin de los parmetros climticos con la base de datos biolgica (especie,
longitud, latitud y elevacin) para evaluar y complementar la informacin disponible sobre la diversidad de mamferos
en la BRTC.
Palabras clave: sustituto de biodiversidad, distribucin geogrca, Mammalia.
Introduction
The analyses of species distribution aimed to identify
priority areas for biodiversity conservation rely mainly on
datasets of point localities where specimens were collected.
Nevertheless, raw data of geographical distributions of
species are frequently incomplete, fragmented and biased
(Reddy and Dvalos, 2003; Rondinini et al., 2006). When
good range data are available for a species or group of
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864 Tellz Valds et al.- Mammals in Tehuacn-Cuicatln domains
species, it is common to use this information as surrogate
of biodiversity patterns in other groups of species where
information is absent or incomplete (Caro and ODoherty,
1999; Pinto et al., 2008). Another choice has been to use
the information contained in patterns at higher levels inthe biological hierarchy, such as vegetation types and
landscape features (Lindenmayer et al., 1991, Margules
and Pressey, 2000). Moreover, with the use of geographic
information systems and interpolators like ANUSPLIN
(Hutchinson, 1991), trustworthy environmental data have
been produced for areas where climatic information was
missing, opening the possibility to use non-biological
elements, such as climatic variables, to explain biological
patterns (Tllez Valds and Dvila Aranda, 2003). Several
studies developed by Hutchinson (1991, 1995a, 1995b,
1997, 1998) and Hutchinson and Gessler (1994) enabled
the 3-dimensional space interpolation of climatic data by
incorporating a digital elevation model to generate rastersurfaces of climatic variables and parameters for Australia.
Using the software package ANUSPLIN, climatic
surfaces of the minimum and maximum temperature and
precipitation, and 19 specic bioclimatic parameters have
recently been derived for Mexico and for the Tehuacn-
Cuicatln valley (Tllez Valds and Dvila Aranda, 2003;
Tllez Valds et al., in press).
The Tehuacn-Cuicatln biosphere reserve (BRTC)
belongs to the oristic province of the Tehuacn-Cuicatln
valley, which is outstanding for its high plant diversity and
endemism (Rzedowski, 1978; Dvila, 1997; Tllez Valds
and Dvila Aranda, 2003). Recently, some surveys and
studies showed that the BRTC is rich in mammal speciesand endemisms compared to other protected natural areas in
Mxico (Botello et al., 2005, 2006a, 2006b; Ramrez Pulido
and Martnez Vzquez, 2007). Nevertheless, studies of
the diversity and distribution of mammals in the BRTC are
limited in the number of localities and species investigated
(Ramrez Pulido and Martnez Vzquez, 2007). Therefore,
current knowledge regarding mammalian distribution
patterns in the BRTC is fragmentary, particularly for
Mexican endemic species (Ramrez Pulido and Martnez
Vzquez, 2007). Thus, the bias and lack of information
prevail for most mammal species in available biological
datasets, and this shortcoming hampered our previous
efforts to model the ecological niche and potential
distribution of mammals within BRTC using BIOCLIM and
GARP. Consequently, the main purpose of this study was
to examine the patterns of mammalian diversity in BRTC
using a climatic domains classication, since domains may
be a good surrogate of biodiversity and may represent the
reserves high biological diversity and endemism rate,
complex topography, and varied environmental conditions
(Tllez Valds and Dvila Aranda, 2003).
Materials and methods
Study area. The Tehuacn-Cuicatln biosphere reserve is
located in southeastern Puebla and northwestern Oaxaca,approximately at 96 55 to 97 44 longitude W and 17
39 to 18 53 latitude N. The reserve has an extension
of about 5 000 km2, but it belongs to a much larger
unit, the oristic province of Tehuacn-Cuicatln valley
(Rzedowski, 1978) which covers about 10 000 km2 and
represents the southernmost semiarid area in Mexico and
in North America. It is one of the most important arid
regions in the world, due to its oristic richness and high
endemism rate (Dvila , 1997; Dvila et al., 2002). Because
the available biological dataset of mammalian distribution
within BRTC was biased and incomplete for most species,
we rather used the more complete and less biased dataset
of mammalian diversity patterns for a region that includedthe Tehuacn-Cuicatln valley and its surrounding areas of
inuence: the Balsas basin, the Mixteca region, the arid
zone of the State of Veracruz, and part of the Sierra Madre
del Sur mountain range.
We dened the climatic domains (geographic units with
similar environments) through multivariate classication
of a data set consisting of climatic estimates for points
on a 1-km grid across our study area. Using ArcView
3.2 (Environmental Systems Research Institute [ESRI],
Redlands, California, U.S.A.), we created a rectangular
polygon with extreme coordinates 19.00o North, 17.00o
South, -96.65o East, and -99.50o West. This polygon
represented our study area and included the Tehuacn-Cuicatln valley and its surrounding areas of inuence: the
Balsas basin, the Mixteca region, the arid zone of the State
of Veracruz, and part of the Sierra Madre del Sur mountain
range. The polygon included an area of approximately
90 000 km2. We overlapped the environmental data with
the biological dataset of point locations (species, longitude,
latitude, and elevation) from mammalian species that fell
in our study area.
Biological dataset. The Comisin Nacional para el
Conocimiento y Uso de la Biodiversidad (CONABIO),
Mxico, provided a dataset of precisely georeferenced
locations from the mammal species that fell in the study
polygon with extreme coordinates 19.00o North, 17.00o
South, -96.65o East, and -99.50o West. We also included
in our dataset georeferenced locations from scats of the
neotropical river otter, (Lontra longicaudis), observed by
Francisco J. Botello (Instituto de Biologa, Universidad
Nacional Autnoma de Mxico), and georeferenced
locations of pictures of mammal species collected by
camera traps during an ecological study of carnivore
mammals in the BRTC (Botello, 2006). With this
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information, and according to the most recent publications
(Villa Ramrez and Cervantes Reza, 2003; Ceballos and
Oliva, 2005; Botello et al., 2005, 2006a, 2006b; Ramrez
Pulido and Martnez Vzquez, 2007), we compiled a list
of mammal species that have been collected, observed, orreported within the BRTC.
Climatic variables and climatic domains classication.
We estimated 19 climate variables (Table 1) for points on
a 1-km grid across our study area (approximately 90,000
grid points) from thin plate smoothing splines (Hutchinson,
1995a) using the ANUSPLIN version 4.3 (Hutchinson and
Gessler, 1994; Hutchinson, 2004) tted to meteorological
station data. The description of the methodology used to
choose and estimate the climate variables, and to classify
the climatic domains has been described fully elsewhere
(Tllez Valds et al., in press). ANUSPLIN has been
successfully used in interpolation exercises (New et
al., 1999, 2002), has good performance in comparativetests of multiple interpolation techniques (Hartkamp et
al., 1999; Jarvis and Stuart, 2001; Hijmans et al., 2005),
and is computationally resourceful and easy to execute.
Interpolation errors were slightly larger than 0.5 Celsius
degrees for maximum and minimum temperatures, and
between 8-13% for monthly mean precipitation (Tllez
Valds et al., in press), which are typical of denser data
networks (Hutchinson, 1997).
Classication results were imported into desktop
geographic information system ArcView 3.2 (ESRI,
Redlands, California, U.S.A.) for inspection of 5-domains
and 10-domains levels of classication, and for their
comparison with digitalized maps of oristic and ecologicalregions, and vegetation types (Rzedowski, 1978). The
climatic domains were overlapped with the biological
database of geographic distribution of mammalian species
in the Tehuacn-Cuicatln valley and surrounding areas of
inuence.
Results
Mammalian dataset. We compiled 9 669 point locations
of mammal species that fell in the study polygon with
extreme coordinates 19.00o North, 17.00o South, -96.65o
East, and -99.50o West. From these point locations,
2 882 were unique registries for 192 species, from which
2 258 point locations belonged to 92 of 98 mammal species
that have been collected, observed or reported within the
BRTC. We listed 98 mammals species that may inhabit
the BRTC, according to the most recent publications (Villa
Ramrez and Cervantes Reza, 2003; Ceballos and Oliva,
2005; Botello et al., 2005, 2006a, 2006b; Ramrez Pulido
and Martnez Vzquez, 2007), and to Ramrez Pulido et al.
(2005) taxonomic classication. The 98 species belonged
to 8 orders, 20 families, and 64 genera of mammals (Table
2). We plotted the georeferenced locations of mammalian
species in Tehuacn-Cuicatln valley and surroundingareas of inuence, as point locations (Figure 1).
We did not obtain georeferenced locations for the vole
Microtus oaxacensis, the big small-eared shrew (Cryptotis
magna), and 4 species of bats: Mexican big-eared bat
(Corynorhinus mexicanus), western red bat (Lasiurus
blossevillii), evening bat (Nycticeius humeralis), and little
yellow bat (Rhogeessa parvula). For the Commissariss
long-tongued bat (Glossophaga commissarisi ) we
compiled just 1 point location. For 7 species we compiled
2 georeferenced locations, and for 19 species we obtained
3 to 9 locations.
Climatic Domain Classication. Noteworthily, at the
5-domains level of classication (Figure 1), domain 4 wasmade up of the Balsas basin, the Tehuacn Valley, and the
Cuicatln Valley. Two main braches for the 5-domains
classication were evident (Figure 2). The rst branch
included the eastern part of the study area, which was the
coastal plain of the Gulf of Mxico, represented by domain
3, and differed in climate from the BRTC and surrounding
areas of inuence, represented by domains 1, 2, 4 and 5.
The second branch showed 2 ramications which separated
domains 1 and 2 from domains 4 and 5, indicating that
the Tehuacn-Cuicatln valley and the Balsas basin had
more similarity with the Sierra Madre del Sur mountain
range, compared to the Sierra Madre Oriental, the Mexican
Transversal Volcanic Belt, and the Sierra Norte de Oaxacamountain range.
At the 10-domains level of classication (Figure 3),
2 main branches showed that climatic differences were
evident and related mainly to precipitation and temperature
rates (Table 3). The rst branch was the group of domains
3, 4, and 6, determined mainly by higher precipitation rates
and less extreme minimum temperature values in the Gulf
of Mexico coastal plain (Figure 4), which differed in climate
from the BRTC and surrounding areas of inuence, and
was made up of tropical rain forests in Veracruz (domain
4), montane forest in Oaxaca and Veracruz (domain 3), and
evergreen tropical forest (domain 6). The second branch
was characterized by temperate and arid environments in
3 ramications: a) domains 2, 10 and 7, b) domains 8 and
9, and c) domains 1 and 5. The rst ramication contained
xeric vegetation such as desert rosette thicket and pine-oak
forest in the Mexican Transverse Volcanic Belt (domain
2), oak and pine-oak forests in the Sierra Madre del Sur
(domain 10), and oak forest in the Sierra Norte de Oaxaca
(domain 7). The second ramication contained deciduous
dry forest, pine forest, oak forest, and pine-oak forest in
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866 Tellz Valds et al.- Mammals in Tehuacn-Cuicatln domains
the Sierra Madre del Sur (domains 8 and 9). The third
ramication contained deciduous dry forest in the Balsas
basin, oak and pine-oak forests in the Sierra Madre del Sur
and Mexican Transversal Volcanic Belt, and thicket in the
Tehuacn-Cuicatln valley. Notably again, the 10-domainsclassication identied climatic similarities between the
Tehuacn-Cuicatln valley and the Balsas basin.
Mammalian diversity in Climatic Domains of BRTC.
Climate domains in the BRTC were characterized
mainly by arid environments, with the addition of some
temperate environments as well. At the 10-domains level
of classication, just 7 of the 10 domains fell in the BRTC
(Table 4), and domain 10, represented by oak and pine-
oak forests, made up 44% of the reserve area. Domain 2
was oak and pine-oak forests, and domains 1 and 5 were
represented by thicket in Tehuacn-Cuicatln valley, and
each of these domains covered over 15 % of BRTC total
area (Figure 3). Domain 7 was represented by oak forestand covered less than 10 % of BRTC area, and domains 3
and 4 made up less than 1 % of the reserves BRTC and
were composed of tropical rain and montane forests (Table
4).
From the total of 98 mammal species that we listed
in the BRTC, georeferenced locations were documented
only in 5 of the 7 domains in the reserve, and the number
of species with point locations varied between 11 and 39
among these 5 domains (Table 4); domains 3 and 4 had no
data and also were poorly represented in BRTC in terms
of area.
The number of species with georeferenced locations
increased substantially for every domain when weconsidered the full information in the study area polygon
(Table 4) with extreme coordinates 19.00o North, 17.00o
South, -96.65o East, and -99.50o West. For example, the
number of species with georeferenced locations in domain
5 increased from 39 to 71. Data behaved similarly for
domains 2, 1, and 10. For domains 3 and 4 that had zero
georeferenced locations of mammal species within the
BRTC, data increased to 53 and 10 species, respectively.
Discussion
Mammalian dataset. The georeferenced locations of
mammalian species in the Tehuacn-Cuicatln valley
and surrounding areas of inuence, plotted in Figure 1
as point locations, showed that the biological dataset was
fragmented, biased, and incomplete. There were several
places without georefenced locations, as well as some
places with higher density of point locations. Although
we listed 98 species, we obtained georeferenced locations
for 92 of them. Also, there were 29 species with few or
very few (1 to 9) point locations. For endemic and rare
species with restricted distribution, the lack of information
may be explained by the difculty of collecting or
observing specimens, as may be the cases of the vole M.oaxacensis and the big small-eared shrew (C. magna),
both endemic to Oaxaca, and the chestnut-bellied shrew
(Sorex ventralis), endemic to Mexico (Ramrez Pulido
et al., 2005; Ramrez Pulido and Martnez Vzquez,
2007). The limited number of species and locations that
have been studied within BRTC are other factors that
may explain the absence of information regarding the
geographic distribution of mammals (Ramrez Pulido
and Martnez Vzquez, 2007). Nevertheless, the absence
of georeferenced locations occurred also for 4 species of
bats that show a wide distribution in Mxico: Mexican big-
eared bat (C. mexicanus), western red bat (L. blossevillii),
evening bat (N. humeralis), and little yellow bat (R.parvula). Similarly, only 1 point location was obtained
for the Commissariss long-tongued bat (G.commissarisi),
and 2 point locations were obtained for other mammal
species without restricted distributions in Mxico: the
silky pocket mouse (Perognathus avus), the tepezcuintle
or lowland paca (Cuniculus paca), and 4 bats: black myotis
(Myotis nigricans), Peales free-tailed bat (Nyctinomops
aurispinosus), big free-tailed bat (Nyctinomops macrotis),
and big crested mastiff bat (Promops centralis).
Our list of mammal species within the BRTC is
appointed to change in the near future, because recent
studies suggest that some species might be extirpated
from the BRTC, and other species may have not beendocumented yet (Botello et al., 2005, 2006a, 2006b;
Ramrez Pulido and Martnez Vzquez, 2007). Ramrez
Pulido and Martnez Vzquez (2007) mention that they
found no evidence to support the assertion of the presence
of jaguarundi (Herpailurus yagouaroundi), puma (Puma
concolor), and jaguar (Panthera onca) within the BRTC,
whereas for the tayra (Eira barbara) they reported that local
people showed them some pelts but they found no further
evidence. Recently, Botello et al. (2005, 2006a, 2006b)
documented the rst photographic registries of margay
(Leopardus wiedii), bobcat (Lynx rufus), and tepezcuintle
(C. paca) in the BRTC, as well as the rst ndings of
scats of neotropical river otter (Lontra longicaudis). In
addition, the American badger (Taxidea taxus) and pocket
gophers (Order Rodentia, Family Geomyidae) are species
acknowledged by local people to be present in the reserve,
but not yet documented with georeferenced data.
Climatic Domains Classication.The climate patterns in our
results revealed that at both the 5-domains and 10-domains
levels of classication, although geographically separated
from each other, the Tehuacn-Cuicatln valley and the
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Table 1. Bioclimatic parameters used for the generation of a climatic domains classication in a region that included the Tehuacn-
Cuicatln valley and its surrounding areas of inuence; with extreme coordinates 19.000North, 17.000 South, -99.500 West and -96.650
East
1. Annual mean temperature (C)
2. Mean temperature diurnal range (C)
3. Isothermality (C)
4. Temperature seasonality (coefcient of variation in %).
5. Maximum temperature of warmest period (C)
6. Minimum temperature of coldest period (C)
7. Temperature annual range (C)
8. Mean temperature of wettest quarter (C)
9. Mean temperature of driest quarter (C)
10. Mean temperature of warmest quarter (C)
11. Mean temperature of coldest quarter (C)
12. Annual precipitation (mm)
13. Precipitation of wettest period (mm)
14. Precipitation of the driest period (mm)
15. Precipitation seasonality (coefcient of variation in %)
16. Precipitation of wettest quarter (mm)
17. Precipitation of driest quarter (mm)
18. Precipitation of warmest quarter (mm)
19. Precipitation of coldest quarter (mm)
Order1 Family1 Number of Species
Artiodactyla Cervidae
Tayassuidae
1
1
Carnivora Canidae
Felidae
Mephitidae
Mustelidae
Procyonidae
2
5
3
2
3
Chiroptera Emballonuridae
Molossidae
Mormoopidae
Phyllostomidae
Vespertilionidae
1
4
3
18
13
Table 2. Taxonomic diversity of mammalian species in Tehuacn-Cuicatln biosphere reserve
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Order1 Family1 Number Of Species
Cingulata Dasypodidae 1
Didelphiomorpha Didelphidae 3
Lagomorpha Leporidae 3
Rodentia Cuniculidae
Heteromyidae
Muridae
Sciuridae
1
3
25
2
Soricomorpha Soricidae 4
Table 2. Continues
1Following the nomenclature classication proposed by Ramrez Pulido et al. (2005).
Table 3. Attributes obtained from 10 domains dened by numerical classication on the basis of climate estimates of approximately
90 000 points on a 1-km grid across Tehuacn-Cuicatln valley and its surrounding areas of inuence
IDAMT
C
MTDR
C
ISO
C
TS
%
MTWP
C
MTCP
C
TAR
C
MTWQ
C
MTDQ
C
MTWQ
C
1 18.4-23.9 12.7-17.2 0.60-0.76 0.42-0.77 28.9-35.6 7.1-13.6 19.5-25.0 19.0-24.6 16.1-23.2 20.3-26.3
2 12.7-20.5 12.7-18.0 0.65-0.74 0.47-0.70 22.5-32.1 2.3-8.9 19.3-25.5 13.2-21.4 10.8-18.2 14.4-22.7
3 15.1-22.2 9.8-14.0 0.58-0.68 0.52-0.74 23.5-33.0 6.8-12.9 16.0-22.2 15.8-23.5 13.7-21.2 16.8-24.6
4 21.5-25.4 10.4-13.9 0.57-0.64 0.70-0.81 30.6-35.7 11.4-16.5 17.9-22.2 22.8-26.9 19.5-24.4 23.8-27.9
5 22.2-28.7 13.1-17.7 0.58-0.71 0.48-0.86 33.9-40.8 9.8-16.6 20.3-27.6 22.7-28.5 19.6-29.4 24.8-31.5
6 19.0-25.3 9.8-12.3 0.57-0.63 0.65-0.79 26.7-34.7 10.7-16.4 16.0-19.6 20.4-26.9 17.8-23.2 21.0-27.8
7 3.4-18.2 6.8-14.1 0.62-0.76 0.20-0.64 8.20-28.1 -0.9-8.9 9.10-20.8 3.4-18.9 3.1-17.4 4.2-20.5
8 21.9-27.7 10.7-15.6 0.66-0.76 0.21-0.45 29.3-37.1 12.9-19.5 14.7-21.6 22.0-27.8 21.3-28.0 22.7-29.0
9 17.2-23.6 10.6-15.4 0.64-0.80 0.21-0.52 26.0-34.1 7.5-14.2 15.0-21.7 17.7-23.9 15.5-23.2 18.6-25.3
10 14.8-20.7 10.4-16.4 0.62-0.75 0.30-0.69 23.2-32.6 4.2-9.7 15.5.-24.2 14.8-21.3 13.4-19.0 16.7-23.2
IDMTCQ
C
AP
%
PWP
mm
PDP
mm
PS
mm
PWQ
mm
PDQ
mm
PHQ
mm
PCQ
mm
1 16.1-22.0 313-1185 20-64 0-0 87-111 160-733 0-59 96-337 6-93
2 10.7-17.9 431-1350 23-69 0-0 80-104 227-791 0-58 146-400 11-61
3 12.9-19.2 1043-2829 53-141 0-11 79-93 594-1504 44-165 183-752 59-235
4 18.5-22.3 979-2697 49-142 0-10 80-100 546-1539 43-156 260-830 53-197
5 19.5-26.4 274-1177 18-64 0-0 90-113 155-730 0-46 86-293 4-62
6 16.2-22.1 2265-4057 107-208 0-16 79-95 1216-2196 126-247 648-1070 129-337
7 2.8-15.8 610-2801 29-123 0-0 77-97 307-1431 0-167 131-483 24-305
8 20.8-26.8 980-2079 56-132 0-0 100-121 629-1302 0-44 224-602 14-50
9 15.5-22.3 836-2082 44-119 0-0 95-113 501-1286 0-53 174-593 18-6410 12.9-18.9 457-1413 29-73 0-0 85-102 236-850 0-50 113-440 12-106
ID = Domain number; AMT = Annual mean temperature; MTDR = Mean temperature diurnal range; ISO = Isothermality; TS =
Temperature Seasonality; MTWP = Mean maximum temperature of warmest period; MTCP = Mean minimum temperature of coldest
period; TAR = Temperature annual range; MTWQ = Mean temperature of wettest quarter; MTDQ = Mean temperature of driest
quarter; MTWQ = Mean temperature of warmest quarter; MTCQ = Mean temperature of coldest quarter; AP= Annual Precipitation;
PWP= Precipitation of wettest period; PDP= Precipitation of driest period; PS= Precipitation Seasonality; PWQ= Precipitation of
wettest quarter; PDQ = Precipitation of driest quarter; PHQ= Precipitation of warmest quarter; PCQ= Precipitation of coldest quarter.
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Table 4. Extension, number of mammalian species, and number of georreferenced locations within 10 domains dened by numerical
classication on the basis of climate estimates of approximately 90 000 points on a 1-km grid across Tehuacn-Cuicatln valley and
its surrounding areas of inuence, a polygon area with extreme coordinates19.00o North, 17.00 o South, -96.65 o East, and -99.50 o West
ID
Study
area
Km2
Studyarea
% area
Total number of
species in study area
Number
of
locations
Number of species collected,
reported or observed for BRTC
Number
of locations
1 17023 18.89 132 580 82 474
2 7915 8.78 71 217 52 190
3 3226 3.58 91 217 53 129
4 3565 3.96 18 30 10 18
5 21144 23.47 102 830 71 754
6 1484 1.65 35 52 16 26
7 5225 5.80 80 305 56 205
8 4497 4.99 69 192 41 130
9 9078 10.08 56 111 41 87
10 16946 18.81 99 34863
272
IDBRTC
Km2BRTC
% area
Number of species collected,
reported or observed for BRTC
Number
of locations
1 1105 18.32 29 51
2 927 15.37 24 32
3 23 0.38 0 0
4 3 0.05 0 0
5 954 15.82 39 136
6 - - - -
7 393 6.52 11 21
8 - - - -
9 - - - -
10 2627 43.55 24 98
ID = Domain number; BRTC = Tehuacn-Cuicatln biosphere reserve.
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870 Tellz Valds et al.- Mammals in Tehuacn-Cuicatln domains
Figure 1. Geographic distribution of 5 domains dened by automatic classication of climatic data for approximately 90 000 points on
a 1-km grid in the Tehuacn-Cuicatln valley and surrounding areas of inuence. Labels indicate the locations of topographic features.
Points represent locations of species of mammals.
Figure 2. Dendrogram describing similarity (climatic distance) between 5 domains dened by automatic classication of climatic data
for approximately 90 000 points on a 1-km grid across the Tehuacn-Cuicatln Valley and surrounding areas of inuence.
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871Revista Mexicana de Biodiversidad 81: 863 - 874, 2010
Figure 3. Geographic distribution of 10 domains dened by automatic classication of climatic data for approximately 90 000 points
on a 1-km grid in the Tehuacn-Cuicatln valley and surrounding areas of inuence. Labels indicate the locations of topographic
features. Points represent locations of species of mammals.
Figure 4. Dendrogram describing similarity (climatic distance) between 10 domains dened by automatic classication of climatic data
for approximately 90 000 points on a 1-km grid across the Tehuacn-Cuicatln valley and surrounding areas of inuence.
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